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Systematic Review of Family Coccidae (Hemiptera:
Coccoidea) in Korean Peninsula with Molecular Phylogeny
밀깍지벌레과(노린재목: 깍지벌레상과)의 계통분류학적 연구
By
Jinyeong Choi
Program in Entomology
Department of Agricultural Biotechnology
Seoul National University
Systematic Review of Family Coccidae (Hemiptera: Coccoidea) in Korean Peninsula with Molecular Phylogeny
Jinyeong Choi Program of Entomology, Department of Agriculture Biotechnology The Graduate School Seoul National University
The systematic studies had three main subjects: i) Taxonomic review of family Coccidae in Korean Peninsula; ii) Molecular phylogeny of family Coccidae; iii) Molecular analyses of genetic diversity and cryptic species of Coccus
hesperidum.
In taxonomic study, a total of thirty-nine species of nineteen genera in the family Coccidae was reviewed with seven new records, Ceroplastes floridensis Comstock, Leptopulvinaria kawaii Tanaka & Amano, Parthenolecanium fletcheri (Cockerell), Pulvinaria hydrangeae Steinweden, P. idesiae Kuwana, P. photiniae Kuwana, and Saissetia Miranda (Cockerell & Parrott). Among previously recorded species, one species, Pulvinaria torreyae, was examined as misidentification. Diagnostic descriptions and identification keys with morphological illustrations and photographs were also given in the present study.
except for Ceroplastinae which is clustered within the major clade of Coccinae. Four tribes of Coccinae are paraphyletic except for Saissetiini in ML tree, especially Coccini and Pulvinariini are irregularly scattered. Also, Paralecanium and
Megapulvinaria, which are sister to Cardiococcinae or Filippiinae, are distinctively
separated from the major clade of Coccinae. Eulecaniinae is paraphyletic in that
Didesmococcus is nested within the clade of Coccinae.
Analyzing the genetic patterns of C. hesperidum revealed high degree of COI haplotype diversity and two cryptic species. In phylogenetic tree based on Maximum Likelihood (ML), all haplotypes of Coccus hesperidum were divided into three distinct clades, which was also supported by a haplotype network. Further, the K2P-distances showed that high genetic divergences among interclades.
Key words: Coccidae, soft scales, systematics, taxonomic review, molecular phylogeny, genetic diversity, cryptic species, the Korean Peninsula
List of tables……….………...…. vii
List of figures……….……….….. ix
PART I. Taxonomic review of the Family Coccidae (Hemiptera: Coccoidea) in the Korean Peninsula Abstract……….………. 1
I. Introduction……….……… 2
1. General introduction of the Family Coccidae……….….. 2
2. Morphology……….………. 3
3. Biology and life cycle……….………. 5
4. Historical review……….………. 8
4.1. Taxonomy and phylogeny of the Family Coccidae………. 8
4.2. History of Korean records……….……… 10
II. Materials and Methods…….………... 14
III. Results………...……….… 16
Family Coccidae Fallén, 1814……….………. 16
Subfamily Ceroplastinae Atkinson, 1886………..………. 17
Genus Ceroplastes Gray, 1828……….……….…… 17
Subfamily Coccinae Fallén, 1814……….………..…… 32
Genus Leptopulvinaria Kanda, 1960……….……….…… 45
Genus Nipponpulvinaria Tanaka, 2008……….………….…… 48
Genus Pulvinaria Tarioni-Tozzetti, 1867……….………... 51
Tribe Saissetiini Hodgson, 1994……….………..…… 70
Genus Parasaissetia Takahashi, 1955………...……... 71
Genus Parthenolecanium Šulc, 1908……….………….……… 75
Genus Saissetia Deplanche, 1859………...……… 88
Subfamily Eriopeltinae Šulc, 1941……….…… 94
Genus Eriopeltis Signoret, 1872……….……… 95
Genus Luzulaspis Cockerell, 1902……….……….… 99
Genus Psilococcus Borchsenius, 1952………….………….…...…… 100
Subfamily Eulecaniinae Koteja, 1988……….………..……… 103
Genus Didesmococcus Borchsenius, 1953……….…..……. 104
Genus Ericerus Guérin-Meneville, 1858……….…………...……….. 105
Genus Eulecanium Cockerell, 1893………...………... 108
Genus Rhodococcus Borchsenius, 1955………...… 112
Genus Sphaerolecanium Šulc, 1908………...…..……… 113
Subfamily Filippiinae Bodenheimer, 1952……….……...………… 116
Genus Metaceronema Takahashi, 1955……….……… 116
Abstract……….………. 127
I. Introduction……….……… 128
II. Materials and methods……….………..…… 129
III. Result……….………...……… 137
IV. Discussion…….……… 142
V. Conclusion……….……….… 148
PART III. Genetic diversity and cryptic species of brown soft scales, Coccus hesperidum (Hemiptera: Coccidae) revealed by molecular analyses Abstract………..……… 149
I. Introduction……….……… 150
II. Materials and methods……….………..… 152
III. Result……….………... 153
IV. Discussion……….……… 155
Conclusion……….……… 162
Literature cited……….……… 163
Appendix I. Living appearances of Korean Coccidae species………….………. 185
Table 2. History of Korean records of the family Coccidae……….…...….. 13 Table 3. Checklist of Coccidae in the Korean peninsula……….…………...……. 126 Table 4. Primers used in the present study……….…. 132 Table 5. Taxa used in this study with GenBank accession numbers…….……..…. 134 Table 6. Haplotype localities with values of frequencies and genetic diversities… 157 Table 7. Percentage of genetic divergence (based on K2P-distances) between clades
of C. hesperidum and two Ceroplastes species for the COI region……. 158 Table 8. AMOVA results of C. hesperidum……….……….. 160 Table 9. Pairwise Fst values between phylogenetic clades of C. hesperidum for COI region………..…………...………... 161
Appendix III. Tables of Biometric Data
Table 10. Ceroplastes ceriferus (Fabricius, 1798) and C. floridensis Comstock, 1881... ……… 213 Table 11. Ceroplastes japonicus Green, 1921…….……….. 214 Table 12. Ceroplastes rubens Maskell, 1893……….………... 215 Table 13. Coccus hesperidum Linnaeus, 1758 and C. pseudomagnoliarum
(Kuwana, 1914)……….……….……….. 216 Table 14. Leptopulvinaria kawaii Tanaka & Amano, 2008 and Nipponpulvinaria horii (Kuwana, 1902) ……….………..… 217
Table 17. Pulvinaria sp. (misreported as Pulvinaria torreyae Takahashi, 1956)… 220 Table 18. Parthenolecanium corni (Bouché, 1844) and P. fletcheri (Cockerell, 1893)..
……… 221 Table 19. Parthenolecanium persicae (Fabricius, 1776)……….…………... 222 Table 20. Saissetia coffeae (Walker, 1852) and Saissetia miranda (Cockerell &
Parrott, 1899)………....………...…. 223 Table 21. Eriopeltis festucae (Fonscolombe, 1834)……….…...……… 224 Table 22. Ericerus pela (Chavannes, 1848)………….………...…… 225 Table 23. Metaceronema japonica (Maskell, 1897) and Takahashia japonica (Cockerell, 1896).………..………..…. 226
Fig. 2. Detail characters of Coccidae……….………….…… 4 Fig. 3. The morphological phylogeny of the family Coccidae, using 105 characters..
……….. 129 Fig. 4. Phylogenetic relationships of the superfamily Coccoidea…….…….…… 130 Fig. 5. Maximum likelihood tree from the combined analysis of all nuclear and mitochondrial fragments. Branches are colored by subfamilies of the family Coccidae. Numbers indicate bootstrap values for each node……...……… 138 Fig. 6. Maximum likelihood tree from the combined analysis of all nuclear and mitochondrial fragments. Branches are colored by tribes of the subfamily Coccinae. Numbers indicate bootstrap values for each node………... 139 Fig. 7. Bayesian inference tree from the combined analysis of all nuclear and mitochondrial fragments. Branches are colored by subfamilies of the family Coccidae. Numbers indicate Bayesian posterior probability values……… 140 Fig. 8. Bayesian inference tree from the combined analysis of all nuclear and mitochondrial fragments. Branches are colored by tribes of the subfamily Coccinae. Numbers indicate Bayesian posterior probability values……… 141 Fig. 9. Maximum likelihood tree for all mtDNA haplotypes of C. hesperidum.…. 158 Fig. 10. Haplotype network for Coccus hesperidum from COI data……...………. 159
Fig. 14. Ceroplastes rubens Maskell, 1893 루비깍지벌레………. 185 Fig. 15. Coccus hesperidum Linnaeus, 1758 무화과깍지벌레……….……..… 186 Fig. 16. Coccus pseudomagnoliarum (Kuwana, 1914) 어리목련깍지벌레…… 186 Fig. 17. Leptopulvinaria kawaii Tanaka & Amano, 2008……….…. 186 Fig. 18. Nipponpulvinaria horii (Kuwana, 1902) 단풍공깍지벌레….………... 186 Fig. 19. Pulvinaria nipponica Lindinger, 1933 무궁화솜깍지벌레……….….. 187 Fig. 20. Pulvinaria hydrangeae Steinweden, 1946……….………….….. 187 Fig. 21. Pulvinaria idesiae Kuwana, 1914……… 187 Fig. 22. Pulvinaria nishigaharae (Kuwana, 1907) 노랑솜깍지벌레……..…… 187 Fig. 23. Pulvinaria photiniae Kuwana, 1914……….……… 188 Fig. 24. Takahashia japonica (Cockerell, 1896) 줄솜깍지벌레……….………. 188 Fig. 25. Parthenolecanium corni (Bouché, 1844) 말채나무공깍지벌레... 188 Fig. 26. Parthenolecanium fletcheri (Cockerell, 1893)………... 188 Fig. 27. Parthenolecanium persicae (Fabricius, 1776) 복숭아공깍지벌레…… 189 Fig. 28. Saissetia coffeae (Walker, 1852) 철모깍지벌레……… 189 Fig. 29. Saissetia miranda (Cockerell & parrott, 1899)………...…. 189 Fig. 30. Didesmococcus koreanus Borchsenius, 1955 진공깍지벌레.……….... 189 Fig. 31. Ericerus pela (Chavannes, 1848) 쥐똥밀깍지벌레…………...……… 190
Appendix II. Illustration of adult females
Fig. 34. Ceroplastes ceriferus (Fabricius, 1798)………...… 191
Fig. 35. Ceroplastes floridensis Comstock, 1881……….………. 192
Fig. 36. Ceroplastes japonicus Green, 1921……….………. 193
Fig. 37. Ceroplastes rubens Maskell, 1893……….…………..… 194
Fig. 38. Coccus hesperidum Linnaeus, 1758……….……… 195
Fig. 39. Coccus pseudomagnoliarum (Kuwana, 1914)……….…….… 196
Fig. 40. Leptopulvinaria kawaii Tanaka & Amano, 2008……….…….… 197
Fig. 41. Nipponpulvinaria horii (Kuwana, 1902)……….……….… 198
Fig. 42. Pulvinaria nipponica Lindinger, 1933……….……… 199
Fig. 43. Pulvinaria hydrangeae Steinweden, 1946……….…………...… 200
Fig. 44. Pulvinaria idesiae Kuwana, 1914……… 201
Fig. 45. Pulvinaria photiniae Kuwana, 1914………....………. 202
Fig. 46. Pulvinaria sp. (misreported as Pulvinaria torreyae Takahashi, 1956)… 203 Fig. 47. Parthenolecanium corni (Bouché, 1844)……….……… 204
Fig. 48. Parthenolecanium fletcheri (Cockerell, 1893)…….……… 205
Fig. 49. Parthenolecanium persicae (Fabricius, 1776)…….……….… 206
Fig. 50. Saissetia coffeae (Walker, 1852)…….………. 207
PART I. Taxonomic review of the Family Coccidae (Hemiptera: Coccoidea) in the Korean Peninsula
Abstract
The family Coccidae is reviewed as 39 species of 19 genera in the Korean Peninsula, including seven new records, Ceroplastes floridensis Comstock, 1881,
Leptopulvinaria kawaii Tanaka & Amano, Parthenolecanium fletcheri (Cockerell), Pulvinaria hydrangeae Steinweden, P. idesiae Kuwana, P. photiniae Kuwana, and Saissetia Miranda (Cockerell & Parrott).
Diagnosis and descriptions for all Korean species are redescribed with keys to subfamilies, tribes, genera, and species. Illustrations, photographs, and other ecological information are also provided.
I. Introduction
1. General introduction of the Family Coccidae
The family Coccidae, also named soft scale insects, is the third largest family of Coccoidea, comprising 1,134 described species of 168 genera in all zoogeographical regions, especially most populated in tropics and subtropics (Hodgson, 1994; Ben-Dov et al., 2015). Taxonomy of this group is primarily based on adult females because of their distinct sexual dimorphism and high accessible to females with sedentary lifestyles (Gullan & Kosztarab, 1997). The female adults are characterized from other families by following characters: well-developed anal plates; deep anal cleft between anal lobes; eversible anal tube; anal ring consist of two sclerotized crescents; a structure of ventral microducts; tarsus without a campaniform sensilla (Hodgson, 1994). Many species are known as important pests in agricultural and ornamental products as well as inhibited pests in quranteen agency, however their faunal study have been poorly conducted. Previously, 32 species of 18 genera are recored in the family Coccidae from the Korean Peninsula (Paek et al., 2010). In addition to the previous species, seven new records, Ceroplastes floridensis Comstock,
Leptopulvinaria kawaii Tanaka & Amano, Parthenolecanium fletcheri (Cockerell), Pulvinaria hydrangeae Steinweden, P. idesiae Kuwana, P. photiniae Kuwana, and Saissetia Miranda (Cockerell & Parrott), are newly reported to the Korean Peninsula,
2. Morphology
The terminology of morphological characters is adapted from Hodgson (1994), Ben-Dov & Hodgson (1997) and Hodgson & Henderson (2000).
3. Biology and life cycle
3.1. Biological characteristics
Sexual dimorphism. Male and female of soft scales have sexually dimorphic life cycles. The females have a sedentary lifestyle and neotenic development including two or three larval stages. Whereas, the males have developed similarly to holometabolism consisting prepupal and pupal stages after two instars. In addition, they have one pair of wings for flight.
Parthenogenesis. Although adult males have an ability to flight, they are fragile and can live for a short time due to lack of functional mouthparts (Gullan & Kosztarab, 1997). As a result, most species are able to reproduce offsprings as parthenogenesis which compensates a low possibility of mating (Hughes-Schrader, 1948). In fact, family Coccidae exhibits diverse types of parthenogenesis like most families of Coccoidea (Nur, 1971; Ross et al, 2010).
3.2. Life cycles of male and female
From 1st to 2nd instar of male & female
1st instar. The first-instar nymphs, also called ‘crawlers’, escape from brood chamber of adult females and play a major role in dispersal. Until this stage, both sexes do not have morphological sexual dimorphism. With well-developed legs, they move to other parts (e.g., branches, twigs and leaves) of existing host plant or try to find new one. After selecting the position, they insert stylet-like mouthparts into plant
cells and feed nutrients from phloem (Hamon & Williams, 1984; Ben-Dov & Hodgson, 1997). During this phase, some species which belong to genus Ceroplates start to secrete soft wax on dorsum and have distinctive wax patterns.
2nd instar. A representative feature of the second-instar is showing sexual dimorphism. During the second-instar nymphs become adults, the differences, such as wax formations and morphological characters, are more remarkable.
From 2nd instar to adult of male
2nd instar male. The second-instar of males are elongate oval shaped and develop secretory organs, especially dorsal tubular ducts in marginal part of body. These organs secrete translucent wax cover and help to fix the nymph to host plant (Ben-Dov & Hodgson, 1997). The presence of dorsal tubular ducts is used for distinguishing males from females in this stage (Hamon & Williams, 1984; Hodgson & Henderson, 2000).
3rd instar male. The third-instar of males are called prepupa stage. During this stage, not only legs and antennae but also wing buds are being distinct. A penial sheath which is a unique character of the male develops at the end of the abdomen. Dorsal and ventral eyes are also present on heads, but they loss functional mouthparts. Therefore, they stop feeding activity after this stage (Ben-Dov & Hodgson, 1997; Hodgson & Henderson, 2000).
4th instar male. Although the forth-instar (i.e., pupa stage) of males have similar morphology to the third-instar, antennae and legs are more segmented and developed. The wing buds are more distinct and the penial sheath is more elongated (Hamon &
Williams, 1984; Ben-Dov & Hodgson, 1997).
Adult male. After last moulting, adult males leave host plants to find mating partners. One of the distinctive features of the adult male is having a pair of fore wings on the mesothorax (hind wings are either absent or reduced as hamulohalteres). Other appendages, such as legs and antennae, are completely developed. Also, they have two or four pair of simple eyes and a pair of ocelli on sclerotized head (Giliomee, 1967; Hodgson & Henderson, 2004).
From 2nd instar to adult of female
2nd and 3rd instar female. The second-instar of females with oval or circular body do not have tubular ducts on dorsum and conspicuous changes in morphology. They only have a change in body size. In case of annual species, the second-instar nymphs which settled on leaves at the first-instar stage move to woody parts of host plants. After resettling on the new places, they overwinter as the second-instar or immature adult stage.
Adult female. After two or three molting, nymphs become adults which have much more dorsal pores and ventral tubular ducts. Most soft scales (e.g., tribe Ceroplastini, Eulecaniini and Saissetiini) gradually make their body swelled and sclerotized. They lay eggs beneath the swelled bodies which act as brood chamber. Some groups (e.g., tribe Coccini) also have the same method like above groups, however their bodies are rather less convex and harden. In case of other groups (e.g., tribe Pulvinariini), they secrete white wax filaments from the ventral glands and lay eggs in the ovisac. Finally, the eggs emerge to crawlers and maintain their life cycles (Ben-Dov & Hodgson, 1997).
4. Historical review
4.1. Taxonomy and phylogeny of the Family Coccidae (Table 1).
Fallén in 1814 introduced the family Coccidae with the type genus Coccus. Targioni Tozzetti (1868) and Signoret (1869) subdivided the Coccidae (i.e. the Coccoidea of today). Targioni Tozzetti (1868) proposed four tribes in Coccidae: Orthezites (Orthezia and relatives), Coccites (the family Pseudococcidae of today), Diaspidites (Diaspididae), and Lecanites (Coccidae). Additionally, the Lecanites contained seven groups including four are related to soft scales which were the Eriophori demum folliculares (including Filippia and Luzulaspis), the Pulcinati (Pulvinaria and Nidulari), the Ceriferi (Ceroplastes, Columnea and Ericerus) and the nudi (Lecanopsis and
Lecanium). Signoret (1869) presented a classification similar to Targioni Tozzetti
(1868), which are four groups: the Diaspides (now regarded as Diaspididae), the Coccides (Pseudococcidae), the Brachyscelides (Eriococcidae) and the Lecanides (Asterolecaniidae, Lecanodiaspididae, Coccidae, Aclerdidae and Tachardiidae). Based on the classification of Signoret (1869), Atkinson (1886) subdivided the Lecanina (=Lecanides) into 5 groups: the Lecaniodiasparia, Signoretiaria (containing
Signoretia (=Luzulaspis), Eriopeltis and Philippia (=Filippia)), Ceroplastaria
(Ceroplastes and Vinsonia), Pulvinariaria (Pulvinaria) and Lecanaria (Lecanium,
Physokermes, Ericerus, Lecanopsis, Aclera and Carteria). These groups are quite similar
to the present classification, except for the last two groups. Handlirsch (1903) used the superfamily Coccoidea grouping all scale insects, whereas Steinweden (1929) restricted
the meaning of the family Coccidae to the soft scale of today. Also, Steinweden (1929) proposed the generic level of classification studying the type species of 32 genera. This classification consisted of three groups: the Coccus (containing Coccus, Eulecanium,
Lecanium, Protopulvinaria, Pulvinaria and Saissetia of today), the Toumeyella
(Neolecanium and Pseudophilippia), and the Exaeretopus (Exaeretopus, Parafairmairia,
Philephedra and Luzulaspis). The other 19 genera were not grouped, however most
presented grouping is used today. Bodenheimer (1953) provided the subfamily groupings including the Ceroplatinae, Coccinae, Eriopeltinae and Filippinae, following the wax structures. Although Borchsenius (1957) agreed to the classification of Bodenheimer, he changed the positon of Eriopeltinae in the Filippiinae, so three groups were presented: Ceroplatinae (with a thick white wax covering the body of adult female), Coccinae (distinct wax absent or present beneath the abdomen of adult female), Filippinae (filamentous wax covering the adult female thoroughly). In addition, the Coccinae contained two tribes, Pulvinariini (wax secreted from beneath the abdomen, enclosing eggs) and Coccini (distinct wax absent and eggs stored under the body. Based on the adult males of 23 species in 19 genera, Gilomee (1967) proposed four genera in the family: the Eulecanium, the Coccus, the Eriopeltis and the Inglisia, This classification was quite different to previous grouping using characters of adult females. Kosztarab & Kozar, 1988 elevated the three generic groups of Gilomee to the subfamilies which were Coccinae, Eulecaniinae, and Eriopeltinae. Tang et al. (1990) and Tang (1991) presented a complex grouping which was quite different to the classification of Giliomee. Based on the female adult, the family divided into four subfamilies: the Pseudopulvinariinae (including Pseudopulvinaria and Mallococcus) and the Filippiinae, Coccinae and
Ceroplastinae. Further, the Filippiinae was subdivided into the tribes Filippiini (consisting of Ceronemina, Ceroplastodina, Eripeltina and Filippiina) and Lecanopsiini (Lecanopsidini); the Coccinae into the tribes Coccini (Coccina and Eulecaniina) and the Pulvinariini (Pulvinariina and Takahashiina); and the Ceroplatinae into the tribes Ceroplastini and Ctenochitonini (Cardiococcina and Ctenochitonina). Most recently, Hodgson (1994) proposed a classification considering the characters of both adult males and females. The Coccidae was divided into 10 subfamilies, including the Coccinae subdivided into four tribes, Coccini, Paralecaniini, Pulvinariini, and Saissetiini.
4.2. History of Korean records (Table 2).
Until now, 32 species of Coccidae have been recored in Korea. In 1928, Machida and Aoyama firstly reported one species, Takahashia japonica (Cockerell), followed by Ceroplastes ceriferus (Fabricius), C. japonicas Green, Nipponpulvinaria horii (Kuwana), Eulecanium kunoense (Kuwana). Additionally, Satio (1931) reported
Ericerus pela (Chavannes), and then Nakayama (1933) described six new records, Coccus hesperidum Linnaeus, Eucalymnatus tessellatus (Signoret), Parasaissetia nigra (Nietner), Parthenolecanium corni (Bouché), Parthenolecanium persicae
(Fabricius), and Saissetia coffeae (Walker). After that, Kanda (1941) reported four species, Eriopeltis festucae (Fonscolombe), Coccus pseudomagnoliarum (Kuwana),
Eulecanium cerasorum (Cockerell), and Parthenolecanium glandi (Kuwana),
followed by Pulvinaria nishigaharae (Kuwana). In 1952, Shiraki reported
Borchsenius studied North Korean species and reported eight species,
Didesmococcus koreanus Borchsenius, Eulecanium kostylevi Borchsenius, E. secretum Borchsenius, Rhodococcus sariuoni Borchsenius, Eriopeltis sachalinensis
Borchsenius, Luzulaspis bisetosa Borchsenius, Parthenolecanium orientale (Borchsenius), Psilococcus ruber Borchsenius from 1955 to 1957. After 1958, Korean researchers reported five species until recently. Paik (1958) described
Metaceronema japonica (Maskell), followed by Pulvinaria torreyae Takahashi, and Eulecanium takachihoi (Kuwana), and then Pulvinaria nipponica Lindinger and P. floccifera (Westwood) reported by Kwon et al. (2005).
Table 1. Historical review of classification of the family Coccidae Steinweden, 1929 Bodenheimer, 1953 Borchsenius,
1957 Giliomee, 1967 Koteja, 1988 Tang et al., 1990 Hodgson, 1994
Coccidae Coccus Toumeyella Exaeretopus Coccidae Ceroplastinae Coccinae Eriopeltinae Filippinae Coccidae Ceroplastinae Coccinae - Pulvinariini - Coccini Filippinae Coccidae Coccus Eulecanium Eriopeltis Inglisia Coccidae Coccinae Eulecaniinae Eriopeltinae Coccidae Coccinae - Coccini - Pulvinariini Ceroplastinae - Ceroplastini - Ctenochitonini Pseudopulvinariinae Filippiinae Coccidae Coccinae - Coccini - Pulvinariini - Paralecaniini - Saissetiini Ceroplastinae Eulecaniinae Filippiinae Eiopeltinae Cardiococcinae Cissococcinae Cyphococcinae Myzolecaniinae Pseudopulvinariinae
Table 2. History of Korean records of the family Coccidae
No. Author Years Recorded species Korean name
1 Machida & Aoyama 1928 Takahashia japonica (Cockerell, 1896) 줄솜깍지벌레 2 Machida & Aoyama 1930 Ceroplastes ceriferus (Fabricius, 1798) 뿔밀깍지벌레
3 Ceroplastes japonicus Green, 1921 거북밀깍지벌레
4 Nipponpulvinaria horii (Kuwana, 1902) 단풍공깍지벌레
5 Eulecanium kunoense (Kuwana, 1907) 공깍지벌레
6 Saito 1931 Ericerus pela (Chavannes, 1848) 쥐똥밀깍지벌레
7 Nakayama 1933 Coccus hesperidum Linnaeus, 1758 무화과깍지벌레
8 Eucalymnatus tessellatus (Signoret, 1873) 남생이깍지벌레
9 Parasaissetia nigra (Nietner, 1861) 검은철모깍지벌레
10 Parthenolecanium corni (Bouché, 1844) 말채나무공깍지벌레
11 Parthenolecanium persicae (Fabricius, 1776) 복숭아공깍지벌레
12 Saissetia coffeae (Walker, 1852) 철모깍지벌레
13 Kanda 1941 Eriopeltis festucae (Fonscolombe, 1834) 사다리털깍지벌레
14 Coccus pseudomagnoliarum (Kuwana, 1914) 어리목련깍지벌레
15 Eulecanium cerasorum (Cockerell, 1900) 포도공깍지벌레
16 Parthenolecanium glandi (Kuwana, 1907) 큰공깍지벌레
17 Kanda 1943 Pulvinaria nishigaharae (Kuwana, 1907) 노랑솜깍지벌레 18 Shiraki 1952 Ceroplastes rubens Maskell, 1893 루비깍지벌레
19 Sphaerolecanium prunastri (Fonscolombe, 1834) 오얏공깍지벌레
20 Borchsenius 1955 Didesmococcus koreanus Borchsenius, 1955 진공깍지벌레
21 Eulecanium kostylevi Borchsenius, 1955 북쪽공깍지벌레
22 Eulecanium secretum Borchsenius, 1955 개야광공깍지벌레
23 Rhodococcus sariuoni Borchsenius, 1955 사리원깍지벌레
24 Borchsenius 1956 Eriopeltis sachalinensis Borchsenius, 1956 진사다리털깍지벌레 25 Borchsenius 1957 Luzulaspis bisetosa Borchsenius, 1952 쌍털깍지벌레
26 Parthenolecanium orientale (Borchsenius, 1957) 애기공깍지벌레
27 Psilococcus ruber Borchsenius, 1952 대륙털깍지벌레
28 Paik 1958 Metaceronema japonica (Maskell, 1897) 과자깍지벌레 29 Paik & Kim 1977 Pulvinaria torreyae Takahashi, 1956 주목솜깍지벌레 30 Paik 1978 Eulecanium takachihoi (Kuwana, 1902) 밤나무깍지벌레 31 Kwon et al. 2005 Pulvinaria nipponica Lindinger, 1933 무궁화솜깍지벌레
II. Materials and Methods
Collection of soft scale insects.
Materials has been collected from all provinces of South Korea, including specimens of National Institute of Agricultural Sciences (NIAS). For taxonomic examinations of soft scale insects, using young adult females are higly recommended because mature adults have swelled and sclerotized bodies which can not be slide mounted. Therefore, it is important to know the periods of occurring young adults as well as their host plants. In the field, living samples would better to be collected together with leaves, stems or twigs of their host plants and transferred to laboratory. After photographing their living appearances, the samples put in 2.0ml tubes with 95% ETOH.
Slide mounted.
In order to exact identifications, the samples which are preserved in alcohol tubes should be slide mounted according to the modified method of Hodgson & Henderson (2000) as follows: Sample preserved in 95% ethy alcohol (ETOH) is transferred to 5ml tube containing 10% potassium-hydroxide (KOH) and then boied for 2-5 hours at 50-60℃ or leave in room temperature for about one or several days. After transferring decolorized specimen to distilled water, pumping the body to extract unnecessary substances using micro tools and leave for 5-10 minutes. Eliminate distilled water and add 1-2 drops of acid fuchsin stain for 50-60 minutes. For
dehydration and cleaning the excess stain, move the specimen to 75% and 95% ETOH each for 5-10 minutes. After that, the sample is transferred into clove oil for over 10 minutes and mounted on a slide glass with Canada balsam.
Examinations.
The slide mounted specimens were studied under a microscope and the digital images were taken by a software (Active measure ver. 3.0.3, Mitani Co. Ltd, Japan) to measure and illustrate for each species. Terminology of morphological characters follows that of Hodgson (1994). All samples examined for the present study are deposited in SNU.
Abbreviations of localities and depositories are as follows:
GG, Gyeonggi-do; GW, Gangwon-do; CB, Chungcheongbuk-do; CN,
Chungcheongnam-do; GB – Gyeongsangbuk-do; GN – Gyeongsangnam-do; JB, Jeollabuk-do; JN, Jeollanam-do; JJ, Jeju-do; NIAS, National Institute of Agricultural Sciences, Suwon, Korea; SNU, Seoul National University, Seoul, Korea.
III. Result
Systematic Accounts
Order Hemiptera 노린재목
Suborder Sternorrhyncha 진딧물아목 Superfamily Coccoidea 깍지벌레상과
Family Coccidae Fallén, 1814 밀깍지벌레과 Type genus: Coccus Linnaeus, 1758
Diagnosis. Well-developed and together quadrate anal plates on anus; a distinct anal cleft between anal lobes; an eversible anal tube to remove honeydew; an anal ring consist of two sclerotized crescents having setae and pores; a structure of ventral microducts; eyespots present on margin; tarsus without a campaniform sensilla (Hodgson, 1994).
Key to Korean subfamilies of Coccidae
1. Ceroplastes-type pores present on dorsum; a sclerotized caudal process present…... ……… Ceroplastinae - Ceroplastes-type pores absent on dorsum; a sclerotized caudal process absent... 2 2. Dorsum covered with a felted ovisac; dorsal tubercles and pocket-like sclerotisations absent; numerous long ventral setae usually present; two stigmatic
spines present in stigmatic area or absent………...………. Eriopeltinae - Dorsum not covered with a felted ovisac; dorsal tubercles and pocket-like sclerotisations occasionally present; numerous long ventral setae absent; zero to three stigmatic spines in each stigmatic clefts………...………. Filippiinae 3. Marginal setae only with simple pointed apices, never bifid or fimbriate; dorsal tubercles and pocket-like sclerotisations never present; each legs without a tibio-tarsal articulatory sclerosis………...… Eulecaniinae - Marginal setae with simple pointed apices, also bifid or fimbriate tips present; dorsal tubercles and pocket-like sclerotisations occasionally present; legs usually with a tibio-tarsal articulatory sclerosis…..………...……….…….……... Coccinae
Subfamily Ceroplastinae Atkinson, 1886 밀깍지벌레아과 Type genus: Coccus Linnaeus, 1758
Diagnosis. A thick wax usually covering entire body; Ceroplastes-type pores present on dorsum; a sclerotized caudal process for lifting anal plates to outside of thick wax; clear areas without pores; distinct characters of stigmatic areas; form of ventral microducts.
Genus Ceroplastes Gray, 1828 밀깍지벌레속 Type species: Ceroplastes janeirensis (Gray, 1828)
Diagnosis. Hemispherical body entirely covered by a thick wax cover; distinctive shape and color of wax; a sclerotized caudal process; only one type of stigmatic spine arranged in stigmatic clefts or margin (Gill, 1988; Hodgson, 1994; Hodgson & Henderson, 2000).
Key to Korean species of Ceroplastes
1. Body covered with reddish brown wax; Ceroplates-type pores present 3 types; ventral tubular duct absent; leg poorly developed………..……… C. rubens - Body covered with white wax; Ceroplates-type pores present 4 types; ventral tubular duct present; legs developed……….……….. 2 2. Body covered with thick white wax having a distinct horn; stigmatic spines conical in triangular shape composed of 5-6 rows……….……….… C. ceriferus - Body covered with turtle-shaped white wax without a distinct projecting horn; stigmatic spines conical, arranged in 2-3 rows………...………. 3 3. Stigmatic spines almost continuously arranged and only 3-5 spinose setae scattered
between anterior and posterior stigmatic clefts……….….... C. japonicus - Stigmaitc spines distinctively separated by continuous 8-9 spinose setae between anterior and posterior stigmatic clefts………..………… C. floridensis
1. Ceroplastes ceriferus (Fabricius, 1798) 뿔밀깍지벌레
Coccus ceriferus Fabricius, 1798: 546. Coccus chilensis Gray, 1828: 7.
Ceroplastes australiae Walker, 1852: 1087. Ceroplastes vayssierei, Mahdihassan, 1933: 561.
Diagnosis. Body covered with thick white wax having a distinct horn; Ceroplates-type pores present 4 Ceroplates-types; stigmatic spines conical in triangular shape composed of 5-6 rows; ventral tubular duct present 1 type; each leg without a tibio-tarsal articulation and an articulation sclerosis.
Living appearance. Body oval, completely covered by thick white wax with a distinct horn and powdery wax bands. Dorsum scarlet to dark red in color, heavily swelled and sclerotized during oviposition. Eggs reddish, deposited in ventral space (Fig. 11).
Slide-mounted material. Body oval to round, 1.2-3.8mm long, 0.9-3.6mm wide, with distinct stigmatic cleft; anal cleft short, present on immature adult but disappeared by caudal process (Fig. 34).
Dorsum. Derm membranous to heavily sclerotized. Dermal areolations absent.
Dorsal tubercles absent. Dorsal setae cylindrical and stout with blunt apices, occasionally slightly pointed, each 5.4-10.0μm long, rather evenly present on dorsum. Dorsal tubular ducts absent. Filamentous ducts with small pores, each 1.5-1.8μm wide, distributed around margin. Ceroplates-type pores present 4 types: Monolocular pores each 4.0-6.0μm wide, rarely present over dorsum. Bilocular pores each 5.6-7.3μm wide, barred bilocular pores evenly scattered on entire dorsum, and irregular bilocular pores mainly distributed around submarginal areas. Trilocular pores each 6.0-8.2μm wide, usually detected over dorsum. Quadrilocular pores each 6.6-8.6μm
wide, present throughout dorsum but some sparsely determined. Preopercular pore absent. Anal plates together elongated heart shape, each with rounded outer angles, 88.4-173.7μm long, 90.5-154.5μm wide. Each plate with 2 apical setae on dorsum (or venter) and 2 discal setae. Anal process elongated in old specimens. Clear areas present in submarginal area of body.
Margin. Marginal setae spinose, usually slightly curved, each 11.3-22.9μm long, with
simple pointed apices, arranged around margin, except for spiracular areas, present about 2-4 laterally between stigmatic area. Stigmatic clefts moderately furrowed each with 40-44 stigmatic spines, conical, bullet-shaped, each 17.0-35.5μm long, arranged in triangular shape composed of 5-6 rows.
Venter. Derm membranous. Multilocular disc-pores 6.8-7.9μm wide, each with
10-12 loculi, mostly with 10 loculi, numerous around vulvar area, less frequently present on anterior area of abdomen and thorax. Spiracular pores 3.2-6.0μm wide, each with 5 loculi, in a band becoming wider from each spiracles and margin. Ventral tubular duct 28.7-39.6μm long, present 1 type with a moderately long outer ductule, a very thin and long inner dutule, occasionally entirely curved, and a quite small terminal gland, scattered around head and anal plates. Ventral microducts small, each 2.3-4.0μm wide, distributed over venter. Ventral submarginal setae sharply pointed, straight, each with a developed basal socket, each 11.3-22.9μm long. Ventral setae acute, straight, each with a rather wide basal socket, each 6.6-12.4μm long, evenly scattered on venter. Leg well developed, each without a tibio-tarsal articulation and a small articulation sclerosis, total length of metathoracic leg each 234.9-368.0μm long: each coxa 89.8-137.0μm long, trochanter+femer 127.7-183.9μm long, tibia+tarsus
146.0-213.0μm long, claw 11.8-20.1μm long. Tarsal digitules thinner and longer than claw digitules. Spiracles very developed, usually posterior peritreme broader than anterior: anterior peritremes each 23.3-90.5μm wide, posterior peritremes each 25.3-92.6μm wide. Antenna 6 segmented, each 130.1-211.9μm long. Clypeolabral shield 117.2-168.7μm wide.
Specimens examined. 5♀, Seoho-dong, Tongyeong-si, GN, 03.viii.2014, coll. J.Y. Choi, on Camellia japonica L. (Theaceae), Coll#. 140803-JY-05; 4♀, Jeodong-ri, Ulleung-eup, Ulleung-gun, GB, 08.viii. 2014, coll. J.Y. Choi, on Camellia japonica L. (Theaceae), Coll#. 140808-JY-01; 1♀, Bokgok-ri, Jipum-myeon, Yeongdeok-gun, GB, 20.x.1999, coll. G.M. Kwon, on Diospyros kaki THUNB. (Ebenaceae), Slide#. M1AV00328; 1♀, Nammunno, Sangdang-gu, Cheongju-si, CB, 21.iii.1997, coll. S.B. An, on Diospyros kaki THUNB. (Ebenaceae), Slide#. M1AV00311; 1♀, Goheung-eup, Goheung-gun, JN, 25.iii.1998, coll. M.L. Lee, on Camellia japonica L. (Theaceae), Slide#. M1AV00286; 1♀, Buchun-ri, Busan-myeon, Jangheung-gun, JN, 18.ix.2001, coll. G.M. Kwon, on Diospyros kaki THUNB. (Ebenaceae), Slide#. M1AV00347.
Host. Aceraceae: Acer japonicus; Amaranthaceae: Amaranthus sp.; Apocynaceae:
Thevetia peruviana; Aquifoliaceae: Ilex aquifolium; I. cornuta; I. crenata; I. japonicus; I. latifolia; I. opaca; I. serrata; I. vomitoria; Araliaceae: Fatsia japonica;
Berberidaceae: Mahonia sp.; Buxaceae: Buxus sempervirens; Celastraceae:
Euonymus europaeus; E. japonicus; Cucurbitaceae: Cucurbita moschata; Ebenaceae: Diospyros kaki; Ericaceae: Azalea sp.; Vaccinium arboreum; Lythraceae: Lagerstroemia indica, Pinaceae: Tsuga canadensis; Pittosporaceae: Pittosporum sp.;
Podocarpaceae: Podocarpus macrophyllus; P. nagi; Rosaceae: Crataegus sp;
Pyracantha coccinea; Spiraea sp.; Rutaceae: Citrus sp.; Sapindaceae: Nephelium lappaceum; Tamaricaceae: Tamarix gallica; Theaceae: Camellia japonica; C. sinensis; Ulmaceae: Ulmus sp.; Verbenaceae: Callicarpa sp. (Gimpel & Davidson,
1974).
Distribution. Cosmopolitan species: Korea, Japan, China, Taiwan, Thailand, Vietnam, Australia, USA, Europe, UK, South America, Africa.
Remarks. Ceroplastes ceriferus is similar to C. pseudoceriferus Green which had been misidentified in Korea (Lee et al., 2012). C. ceriferus is differentiated from C.
pseudoceriferus by 15 marginal setae between anterior stigmatic clefts (about 40 in
C. pseudoceriferus) and multilocular disc-pores absent around procoxa (Hodgson & Peronti, 2012). Although C. pseudoceriferus was not confimed in this study, it is possible to exist in Korea.
2. Ceroplastes floridensis Comstock, 1881
Ceroplastes floridensis Comstock, 1881: 331.
Diagnosis. Body covered with thick and bright white wax without a distinct projecting horn; Ceroplates-type pores present 4 types; stigmatic spines conical, arranged in 2-3 rows; ventral tubular duct present 1 type; each leg without a tibio-tarsal articulation and an articulation sclerosis.
Living appearance. Body oval, thoroughly coated with bright white wax with powdery wax bands on margin. Wax surface rather flat, without a distinct projecting
horn. Dorsum reddish, highly swelled and sclerotized during oviposition. Eggs reddish, present under venter (Fig. 12)
Slide-mounted material. Body oval, 1.3-1.9mm long, 0.9-1.3mm wide, with distinct stigmatic cleft; anal cleft rather short, present on young adult, but disappeared by caudal process (Fig. 35).
Dorsum. Derm membranous to heavily sclerotized. Dermal areolations absent.
Dorsal tubercles absent. Dorsal setae very short with bluntly pointed apices, each 3.4-4.3mm long, scarcely distributed over dorsum. Dorsal tubular ducts absent. Filamentous ducts not detected. Ceroplates-type pores present 4 types: Monolocular pores each 2.8-4.0μm wide; Bilocular pores each 3.9-4.5μm wide, present 2 types, barred and irregular bilocular pores; Trilocular pores each 3.7-6.0μm; Quadrilocular pores each 3.4-4.0μm, most pores scattered on entire dorsum and intermixed with each pore. Preopercular pore absent. Anal plates each with rounded outer angles, 99.3-126.1μm long, 94.2-124.8μm wide. Each plate with 4 apical setae setae. Anal process not observed. Clear areas present in submarginal area of body.
Margin. Marginal setae spinose, mostly straight or slightly curved, each 16.8-20.0μm
long, with simple pointed apices, arranged around margin, except for spiracular areas, about 41-44 anteriorly between anterior stigmatic areas. Stigmatic clefts moderately furrowed each with 30-44 stigmatic spines. Stigmatic spines conical, bullet-shaped, each with a narrow basal socket, each 9.9-14.1μm long, arranged in 2-3 rows, distinctively separated by continuous 8-9 spinose setae between anterior and posterior stigmatic clefts.
loculi, frequently present around vulvar area, also sparsely scattered on anterior area of abdomen and thorax. Spiracular pores 3.4-3.9μm, each with 5 loculi in 3-4 pores wide between each spiracles and margin. Ventral tubular duct 18.9-21.3μm long, present 1 type with a developed outer ductule, rather broad inner ductule and a swelled terminal gland, usually present around submargin. Ventral microducts small, each 1.1-2.6μm wide, evenly distributed over venter. Ventral submarginal setae sharply spinose, each 6.8-8.1μm long. Ventral setae with 2 pairs of long and short setae between antennae and other setae acute, straight, each 5.0-8.4μm long, rarely present on entire venter. Leg well developed, each without a tibio-tarsal articulation and an articulation sclerosis, total length of metathoracic leg each 351.2-524.0μm long: each coxa 86.1-137.2μm long, trochanter+femer 118.0-173.2μm long, tibia+tarsus 130.2-199.7μm long, claw 11.1-14.5μm long. Tarsal digitules thinner and longer than claw digitules. Spiracles developed, mostly posterior peritreme broader than anterior: anterior peritremes each 20.3-39.3μm wide, posterior peritremes each 21.6-45.0μm wide. Antenna 6 segmented, each 201.3-275.3μm long. Clypeolabral shield 109.1-152.7μm wide.
Specimens examined. 2♀, Yanghwa-dong, Yeongdeungpo-gu, Seoul, 02.viii.2014, coll. J.Y. Choi, on Poaceae sp., Coll#. 140802-JY-06; 2♀, same locality, date and collector, on Ternstroemia japonica (Wight & Arn.) (Theaceae), Coll#. 140802-JY-09; 1♀, Seoho-dong, Tongyeong-si, 03.viii.2014, GN, coll. S.H. Lee, Coll#. 140803-JY-03; 1♀, Jeodong-ri, Ulleung-eup, Ulleung-gun, GB, 08.viii.2014, coll. Y.L. Lee, on Machilus thunbergii SIEB. et ZUCC. (Lauraceae), Coll#. 140808-JY-01; 1♀,
Jang-dong, Dong-gu, Gwangju, 01.i.2016, coll. J.Y. Choi, on Euonymus japonicus Thunb. (Celastraceae), Coll#. 160108-JY-01.
Host. Anacardiaceae: Mangifera indica; Schinus terebinthifolius; Aquifoliaceae: Ilex
cornuta; I. crenata; I. vomitoria; Araliaceae: Aralia sp.; Hedera helix; Sciadophyllum
sp.; Burseraceae: Bursera simaruba; Celastraceae: Euonymus japonicas; Ebenaceae:
Diospyros kaki; Ericaceae: Pernettya sp.; Vaccinium sp.; Lauraceae: Lindera benzoin, Persea borbonia; Myrtaceae: Eugenia compacta; Eugenia jambolana; Psidium guajava; P. pomiferum; Pinaceae: Pinus elliotti; Tsuga canadensis; Platanaceae: Platanus sp.; Poaceae sp.; Polygonaceae: Coccoloba diversifolia; Punicaceae: Punica granatum; Rosaceae: Prunus persica; P. salicina; Pyrus communis; P. cydonia; P. malus; Rubiaceae: Gardenia jasminoides; Serraceniaceae: Serracenia minor (Hall,
1922; Gimpel et al., 1974; Hamon & Williams, 1984)
Distribution. Korea, Japan, China, Taiwan, Vietnam, Indonesia, Australia, USA, Canada, Europe, South America, Africa.
Remarks. Because of morphological similarities, Ceroplastes floridensis may have been overlooked as C. japonicas Green in Korea. However, both species are easily distinguished by arrangements of stigmatic spines (See above the key to species of
Ceroplastes). In this study, C. floridensis is newly reported from Korea.
3. Ceroplastes japonicus Green, 1921 거북밀깍지벌레
Diagnosis. Body covered with turtle-shaped and yellowish white wax without a distinct projecting horn; Ceroplates-type pores present 4 types; stigmatic spines conical, arranged in 2-3 rows; ventral tubular duct present 1 type; each leg without a tibio-tarsal articulation and an articulation sclerosis.
Living appearance. Body oval, thoroughly coated with turtle-shaped white wax with powdery wax bands on margin. Wax surface flat or rough, without a distinct projecting horn. Dorsum reddish dark brown in color, strongly swelled and sclerotized during oviposition. Eggs reddish, stored under venter (Fig. 13).
Slide-mounted material. Body oval to circular, 1.7-3.5mm long, 1.1-3.1mm wide, with distinct stigmatic cleft; anal cleft very short, present on young adult but obscured by caudal process (Fig. 36).
Dorsum. Derm membranous to strongly sclerotized. Dermal areolations absent.
Dorsal tubercles absent. Dorsal setae rather broad with bluntly pointed apices, each 5.1-8.3mm long, rarely scattered over dorsum. Dorsal tubular ducts absent. Filamentous ducts not detected. Ceroplates-type pores present 4 types: Monolocular pores each 3.9-5.3μm wide; Bilocular pores each 4.1-5.3μm wide, present 2 types, barred and irregular bilocular pores; Trilocular pores each 4.0-7.0μm; Quadrilocular pores each 4.0-6.6μm, most pores present on entire dorsum and intermixed with each pore. Preopercular pore absent. Anal plates each with rounded outer angles, 138.6-160.4μm long, 109.9-137.0μm wide. Each plate with 4 apical setae. Anal process elongated in matured adult. Clear areas present in submarginal area of body.
Margin. Marginal setae spinose, mostly straight or slightly curved, each 12.2-30.3μm
areas, about 38-40 anteriorly between anterior stigmatic areas. Stigmatic clefts moderately deep each with 98-123 stigmatic spines on each side. Stigmatic spines conical, bullet-shaped, each with a narrow basal socket, each 19.5-27.8μm long, arranged in 2-3 rows, continuously arranged between anterior and posterior stigmatic clefts. Only 3-5 spinose setae scattered between each stigmatic clefts.
Venter. Derm membranous. Multilocular disc-pores 5.3-7.8μm wide, each with 10
loculi, uncounted pores scattered around vulvar area, also rarely present on anterior area of abdomen and thorax. Spiracular pores 3.4-5.7μm, each with 5 loculi in a broad band between each spiracles and margin. Ventral tubular duct 21.2-41.4μm long, present 1 type with a developed outer ductule, a greatly or moderately broad inner ductule, and a swelled terminal gland, mainly distributed around submargin. Ventral microducts small, each 1.6-3.6μm wide, evenly present over venter. Ventral submarginal setae acute, straight or slightly curved, each 7.0-12.1μm long. Ventral setae with 2 pairs of long and short setae between antennae and other setae sharply pointed, straight, each 4.4-9.6μm long, scarcely scattered on entire venter, but frequently detected around anal plates. Leg well developed, each without a tibio-tarsal articulation and a small articulation sclerosis, total length of metathoracic leg each 543.2-635.7μm long: each coxa 131.3-160.2μm long, trochanter+femer 182.7-222.7μm long, tibia+tarsus 200.3-230.0μm long, claw 15.7-23.2μm long. Tarsal digitules thinner and longer than claw digitules. Spiracles well developed, mostly posterior peritreme broader than anterior: anterior peritremes each 41.9-53.4μm wide, posterior peritremes each 50.8-60.8μm wide. Antenna 6 segmented, each 285.3-327.7μm long. Clypeolabral shield 141.8-163.1μm wide.
Specimens examined. 2♀, Buksan-ri, Seoun-myeon, Anseong-si, GG, 29.iv.1999, coll. S.H. Lee, on Diospyros kaki THUNB. (Ebenaceae), Slide#. M1AV00078, M1AV00079; 1♀, Sangmaengbang-ri, Geundeok-myeon, Samcheok-si, GW, 29.v.1999, coll. J.Y. Choi, on Diospyros kaki THUNB. (Ebenaceae), Slide#. M1AV00082; 3♀, Cheonji-dong, Seogwipo-si, JJ, 14.ix.2014, coll. J.Y. Choi, on
Pittosporum tobira (Thunb.) (Pittosporaceae), Coll#. 140914-JY-01; 1♀, Suwon-ri,
Hallim-eup, Jeju-si, JJ, 17.ix.1996, coll. S.B. An, on same host plant, Slide#. M1AV00121; 3♀, same locality, 18.ix.1996, coll. and host data absent, Slide#. M1AV00186, M1AV00184, M1AV00185.
Host. Apocynaceae: Nerium oleander; Trachelospermum asiaticum; Aquifoliaceae:
Ilex integra; Berberidaceae: Berberis sp.; Epimedium colchicum; Buxaceae: Buxus sempervirens; Celastraceae: Euonymus japonicus; Cornaceae: Cornus mas; Svida sp.;
Cycadaceae: Cycas revoluta; Ebenaceae: Diospyros kaki; Ehretiaceae: Ehretia
acuminata; Elaeagnaceae: Elaeagnus pungens; Elaeocarpaceae: Elaeocarpus decipiens; Lauraceae: Laurus nobilis; Machilus thunbergii; Magnoliaceae: Magnolia grandiflora; Moraceae: Morus sp.; Myrtus communis; Myrtaceae: Feijoa sellowiana;
Pittosporaceae: Pittosporum tobira; Podocarpaceae: Podocarpus nagi; Rosaceae:
Cerasus avium; C. vulgaris; Crataegus sp.; Cydonia vulgaris; Eriobotrya japonica; Malus domestica; Persica vulgaris; Prunus laurocerasus; P. yedoensis; P. sinensis;
Rutaceae: Citrus sp.; Poncirus trifoliata; Salicaceae: Salix glandulosa; S. saidaeana; Theaceae: Camellia japonica; Eurya japonica; Thea sinensis (Tahakashi,1956; Borchsenius, 1957).
4. Ceroplastes rubens Maskell, 1893 루비깍지벌레
Ceroplastes rubens Maskell, 1893: 214. Ceroplastes rubensminor Maskell, 1897b: 309.
Diagnosis. Body covered with reddish brown wax without a distinct projecting horn;
Ceroplates-type pores present 3 types; stigmatic clefts extremely deep, each with
22-26 stigmatic spines; ventral tubular duct absent; legs poorly developed.
Living appearance. Body oval, entirely concealed by moderately thick wax reddish brown wax with distinct curved wax bands on margin. Wax surface flat or roundly convex, without a distinct projecting horn. Dorsum dark red in color, considerably swelled and sclerotized during oviposition. Eggs reddish, laid in swollen venter (Fig. 14).
Slide-mounted material. Body oval to round, 1.7-3.8mm long, 1.3-3.2mm wide, with extremely furrowed stigmatic cleft; anal cleft rather short, present on young adult but shrunk by caudal process (Fig. 37).
Dorsum. Derm membranous to severely sclerotized. Dermal areolations absent.
Dorsal tubercles absent. Dorsal setae stout and short each with a developed basal socket and a quite blunt apex, each 4.4-6.4mm long, evenly scattered over dorsum. Dorsal tubular ducts absent.
Filamentous ducts absent. Ceroplates-type pores present 3 types: Monolocular pores each 4.0-5.3μm wide; Bilocular pores each 4.8-7.2μm wide, barred bilocular pores,
present 2 types, barred and irregular bilocular pores; Trilocular pores each 6.1-9.0μm wide, irregular trilocular pores, most pores present on entire dorsum and intermixed with each pore, but Quadrilocular pores absent. Preopercular pore absent. Anal plates each with rather rounded outer angles, 140.8-160.3μm long, 117.8-149.2μm wide. Each plate with 4 apical setae. Anal process proceeded slightly in matured adult. Clear areas about 10 present in submedian area of body.
Margin. Marginal setae similar to dorsal setae, but slightly longer and less blunt than
the latter, each 4.5-7.6μm long, arranged around margin, except for spiracular areas, present about 2-4 laterally between stigmatic areas. Stigmatic clefts extremely deep each with 22-26 stigmatic spines: Median setae conical, mostly with blunt and slightly curved apices, each 39.9-66.6μm long. Lateral setae hemispherical with roundly convex apices, each 26.1-35.6μm long. The others similar to lateral setae, but much smaller than the latter, each 10.0-16.1μm long, present on ventral stigmatic clefts.
Venter. Derm membranous. Multilocular disc-pores 6.3-8.6μm wide, each with 10
loculi, many pores present around vulvar area, also rarely scattered on anterior area of abdomen and thorax. Spiracular pores 5.0-6.6μm wide, each with 5 or 6 loculi, usually with 5 loculi, arranged in a broad band between each spiracles and margin. Ventral tubular duct absent. Ventral microducts small, each 3.1-4.6μm wide, frequently scattered over venter. Ventral submarginal setae spinose, straight and rather stout, each 5.9-9.6μm long. Ventral setae with 2 pairs of long setae between antennae and other setae sharply pointed, usually straight, each 7.7-9.4μm long, frequently distributed on entire venter. Leg very short and poorly developed, total
length of metathoracic leg each 156.2-212.8μm long: each coxa 56.2-71.9μm long, trochanter+femer 37.0-64.6μm long, tibia+tarsus 47.7-73.6μm long, claw 6.4-12.9μm long. Tarsal digitules longer than claw digitules. Spiracles well developed, posterior peritreme slightly broader than anterior: anterior peritremes each 55.6-70.6μm wide, posterior peritremes each 55.6-75.3μm wide. Antenna 6 segmented, each 155.9-199.4μm long. Clypeolabral shield 151.5-181.1μm wide.
Specimens examined. 2♀, Nohyeong-dong, Jeju-si, JJ, 13.ix.2014, coll. J.Y. Choi, on Citrus sp. (Rutaceae), Coll#. 140913-JY-09; 2♀, Gunnae-ri, eup, Wando-gun, JN, 04.vii.2014, same collector, on Camellia japonica L. (Theaceae), Coll#. 140704-JY-06; 2♀, Seogwipo-si, JJ, 18.i.1972, collector data absent, on Euonymus
alatus (Thunb.) (Celastraceae), Slide#. M1AV00377, M1AV00378; 2♀,
Topyeong-dong, Seogwipo-si, JJ, 5.ix.2001, coll. G.M. Kwon, on Citrus unshiu Marcovitch (Rutaceae), Slide#. M1AV00396, M1AV00399; 2♀, Donghong-dong, Seogwipo-si, JJ, 27.xi.1996, coll. K.S. Lee, on Ligustrum obtusifolium Sieb. & Zucc. (Oleaceae),
Slide#. M1AV00408, M1AV00412.
Host. Acanthaceae: Acer palmatum; Apocynaceae: Alyxia olivaeformis; Aquifoliaceae: Ilex cornuta; I. latifolia; I. oldhami; I. serrata; Araceae: Aglaonema
pictum; Anthurium andraeanum; Araliaceae: Dizygotheca elegantissima; Hedera helix; Araliceae: Philodendron gigantium; Aspleniaceae: Asplenium nidum;
Asteraceae: Helianthus sp.; Buxaceae: Buxus microphylla; Caprifoliaceae: Viburnum sp.; Clusiaceae: Calophyllum inophyllum; Celastraceae: Euonymus alatus; Cycadaceae: Cycas sp.; Dicksoniaceae: Cibotium sp.; Ebenaceae: Diospyros kaki; Fabaceae: Acacia sp.; Lecythidaceae: Barringtonia racemosa; Loranthaceae:
Loranthus sp.; Magnoliaceae: Magnolia salicifolia; Moraceae: Cudrania javanesis;
Myrtaceae: Metrosideros collina; Psidium guajava; Rhodomyrtus tomentosa; Oleaceae: Ligustrum obtusifolium; Orchidaceae: Grammatophyllum sp.; Pinaceae:
Pinus parviflora; Polypodiaceae: Polypodium sp.; Rhizophoraceae: Rhizophora sp.;
Rosaceae: Chaenomeles sp.; Malus sp.; Rubiaceae: Gardenia jasminoides; Rutaceae:
Citrus deliciosa; C. reticulata; C. unshiu; Sapindaceae: Litchi sp.; Sapotaceae: Calocarpum sp.; Schisandraceae: Kadsura japonica; Sinopteridaceae: Pellaea sp.;
Taxaceae: Cephalotaxus sp.; Theaceae: Camellia japonica; C. rusticans, C. sasanqua; Ulmaceae: Celtis sp.; Zingiberaceae: Alpinia purpurata (Gimpel & Davidson, 1974). Distribution. Cosmopolitan species: Korea, Japan, China, Taiwan, Thailand, Vietnam, Indonesia, Australia, USA, Africa.
Remarks. Ceroplastes rubens is easily recognized by wax test of reddish brown.
Subfamily Coccinae Fallén, 1814 무화과깍지벌레아과
The subfamily Coccinaewas divided into about 55 genera belonging to four tribes (Hodgson, 1994).
Key to Korean tribe of Coccinae
1. Body with white wooly ovisac under abdomen; three or four types (rarely two) of ventral tubular ducts present……….………....…………. Pulvinariini - Body without white wooly ovisac under abdomen; one or two types of ventral
2. Ventral tubular ducts present on medial area of thorax or absent; multilocular disc pores only present around vulvar area, not on medial thorax and head... Coccini - Ventral tubular ducts present on submarginal area; multilocular disc pores present around vulvar area, also medial thorax………...………….. Saissetiini
Tribe Coccini Fallén, 1814 무화과깍지벌레족 Type genus: Coccus Linnaeus, 1758
Diagnosis. Ventral tubular ducts restricted to medial area of thorax or absent; absence of dorsal tubular duct except for some species in Coccus; dorsum without pocket-like sclerotizations; eyespots closely located on margin; differentiated stigmatic spines; unsclerotized stigmatic areas; multilocular disc pores only present around vulvar area, not on medial thorax and head (Hodgson, 1994).
Key to Korean genera of Coccini
1. Irregular polygonal plates present on dorsum………..……..…. Eucalymnatus - Irregular polygonal plates absent on dorsum……….…….. Coccus
Genus Coccus Linnaeus, 1758 무화과깍지벌레속 Type species: Coccus hesperidum Linnaeus, 1758
area; legs with a tibio-tarsal articulation sclerosis; blunt or cylindrical dorsal setae; marginal setae with bifid or fimbriate apices; each triangular shaped anal plates with a few apical setae (Hodgson, 1994).
Key to Korean species of Coccus
1. Dorsal tubercles present; Marginal setae with bifid or fimbriate apices, occasionally with pointed tips; Antenna 7 segmented……….……... C. hesperidum - Dorsal tubercles absent; Marginal setae with pointed apices; Antenna 8 Segmented………... C. pseudomagnoliarum
5. Coccus hesperidum Linnaeus, 1758 무화과깍지벌레
Coccus hesperidum Linnaeus, 1758: 455. Calypticus laevis Costa, 1829: 11. Coccus patellaeformis Curtis, 1843: 517. Chermes lauri Boisduval, 1867: 340. Lecanium angustatus Signoret, 1873b: 398. Kermes aurantj Alfonso, 1875: 431. Lecanium alienum Douglas, 1886: 77. Lecanium minimum Newstead, 1892: 141.
Lecanium assimile amaryllids Cockerell, 1893b: 53. Lecanium terminaliae Cockerell, 1893c: 254. Lecanium ceratoniae Gennadius, 1895, cclxxvii.
Lecanium nanum Cockerell, 1896b.
Lecanium minimum pinicola Maskell, 1897b: 310. Lecanium flaveolum Cockerell, 1897b: 52.
Lecanium ventrale Ehrhorn, 1898: 245.
Lecanium (Calymnatus) hesperidum pacificum Kuwana, 1902a: 30. Lecanium signiferum Green, 1904: 197.
Coccus hemisphaerides Lindinger, 1929: 109-110. Coccus jungi Chen, 1936: 218.
Diagnosis. Dorsal tubercles present; dorsal setae cylindrical and stout, with blunt apices; dorsal tubular ducts present or not; marginal setae with pointed, bifid or fimbriate apices; multilocular disc-pores each with 10 loculi; leg each with a tibio-tarsal articulatory sclerosis; antenna 7 segmented.
Living appearance. Body elongate oval to round, occasionally diverse asymmetrical shapes depend on host plants, and relatively flat or moderately convex during oviposition. Dorsum greenish or yellowish brown in color, usually with irregular black or brown spots (Fig. 15).
Slide-mounted material. Body elongate oval to circular, 2.0-3.8mm long, 1.6-2.8mm wide, with rather distinct stigmatic cleft; anal cleft approximately 1/6-1/7 of body length (Fig. 38).
Dorsum. Derm membranous. Dermal s developed and small, each with a
microductule. Dorsal tubercles convex, each with an inner filamentous ductule, present on submarginal area, 4-6 in total on each side: 1 or 2 between apex of head
and anterior stigmatic cleft, 1 between anterior and posterior stigmatic clefts and 2 or 3 between posterior stigmatic cleft and anal cleft. Dorsal setae cylindrical and stout with blunt apices, each 5.5-9.4μm long, evenly scattered on dorsum. Dorsal tubular ducts not detected. Preopercular pore round to oval, distributed in a dense group in front of anal plates. Anal plates each triangular in shape, 135.2-175.4μm long, 110.9-152.7μm wide, mostly posterolateral margin rather longer than anterolateral margin: anterolateral margin 86.2-109.7μm long, posterolateral margin 91.7-122.5μm long. Each plate with 2 inner margin setae, 1 outer margin setae, and 1 apical setae on dorsum (or venter).
Margin. Marginal setae spinose, straight or slighly curved, each 18.5-36.6μm long,
usually with bifid or fimbriate apices, also simple pointed tips frequently detected according to specimens, present about 8-10 laterally between stigmatic areas. Stigmatic clefts rather distinct each with 3 stigmatic spines, median spine slightly curved, and more or less 2 times as long as lateral spine: medians 38.2-73.8μm long, laterals 4.9-44.2μm long.
Venter. Derm membranous. Pregential disc-pores 3.2-4.7μm wide, mostly each with
10 loculi, relatively a small number present around vulvar area. Spiracular pores 3.6-4.5μm wide, each with 5 loculi, in a narrow or single band between each spiracles and margin. Ventral tubular duct present 1 type with a moderately narrow inner ductule and a developed flower-head-like terminal gland, present around each coxa of legs and anal plates. Ventral microducts quite small, rarely distributed over entire venter. Ventral setae with 3 pairs of long pregenital setae present, also about 2 pairs of long setae between antennae and other setae sharply spinose, scarcely present on
entire venter. Leg well developed, each with a tibio-tarsal articulation and a small articulation sclerosis, total length of metathoracic leg each 273.8-548.6μm long: each coxa 89.8-137.0μm long, trochanter+femer 127.7-183.9μm long, tibia+tarsus 146.0-213.0μm long, claw 11.8-20.1μm long. Tarsal digitules thinner and longer than claw digitules. Spiracles normally developed, mostly posterior peritreme broader than anterior: anterior peritremes each 22.4-36.1μm wide, posterior peritremes each 28.0-49.2μm wide. Antenna 7 segmented, each 237.8-329.1μm long. Clypeolabral shield 102.2-142.5μm wide.
Specimens examined. 2♀, Punggok-ri, Gochon-eup, Gimpo-si, GG, 20.v.1998, coll. G.M. Kwon, on Ficus benjamina L. (Moraceae), Slide#. M1AV00453, M1AV00456; 2♀, Neunggok-dong, Siheung-si, GG, 20.V.1998, same collector and host, Slide#. M1AV00449, M1AV00452; 2♀, Sucheong-dong, Osan-si, GG, 06.vi.2015, coll. J.Y. Choi, on Schefflera actinophylla (Endl.) (Araliaceae), Coll#. 150601-JY-04; 2♀, Cungjeongno, Jung-gu, Seoul, 25.iii.2015, same collector, on Dracaena sp. (Asparagaceae), Coll#. 150325-JY-01; 2♀, Cheolli-dong, Andong-si, GB, 07.vi.2015, same collector, on Schefflera sp. (Araliaceae), Coll#. 150607-JY-06; 2♀, Cheonjeon-ri, Sinbuk-eup, Chuncheon-si, GW, 31.v.2015, same collector, on Heteropanax sp. (Araliaceae), Coll#. 150531-JY-01; 2♀, Sincheon-dong, Dong-gu, Daegu, 06.vi.2015, same collector and host, Coll#. 150606-JY-12; 2♀, Gayang-dong, Dong-gu, Daejeon, 06.vi.2015, same collector and host, Coll#. 150606-JY-03; 1♀, Sacheon-ri, Deoksan-myeon, Yesan-gun, CN, 01.vi.2015, same collector, on Schefflera actinophylla (Endl.) (Araliaceae), Coll#. 150424-JY-01; 1♀, Yeon-dong, Jeju-si, JJ, 17.iv.2003, coll. G.M. Kwon, on Schefflera sp. (Araliaceae), Slide#. M1AV00429; 1♀, Sinhyo-dong,
